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Specimen of the Recently Described Calliophis Salitan (Squamata: Elapidae), with the First Report of the Species from Mindanao Island, Southern Philippines

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Specimen of the Recently Described Calliophis Salitan (Squamata: Elapidae), with the First Report of the Species from Mindanao Island, Southern Philippines Herpetology Notes, volume 14: 1027-1035 (2021) (published online on 03 August 2021) On the second (or, rather, the first) specimen of the recently described Calliophis salitan (Squamata: Elapidae), with the first report of the species from Mindanao Island, southern Philippines Rafe M. Brown1,*, Marta Calvo-Revuelta2, Johana Goyes Vallejos3, Alberto Sánchez-Vialas2, and Ignacio De la Riva2 Abstract. We report on the second known specimen of the recently described banded Asian coralsnake, Calliophis salitan, identified among the Museo Nacional de Ciencias Naturales collections of Dr. Hipólito Fernández, made in 1886. The specimen was collected on Mindanao Island in the southern Philippines as part of the “Comisión Central de Manila,” the Spanish Crown’s effort to catalogue and showcase Philippine biodiversity for the Madrid public. We assign this important specimen to C. salitan (formerly known from a single specimen collected on Dinagat Island) based on its highly distinctive external phenotypic characteristics (scalation, body size, and inferred colour pattern). The discovery of C. salitan on Mindanao significantly extends the known geographic range of this species and partially fills an intermediate distributional hiatus – a large geographic gap in the distribution of large-bodied, long-glanded coralsnakes. This report partially resolves a biogeographic enigma surrounding the question of how a distinctive evolutionary lineage of elapid snakes – more closely related to the Calliophis bivirgatus group from Borneo and other landmasses of Sundaland than to any other Philippine elapids – could have colonized only a small island (Dinagat) in the geographic interior of the archipelago. The answer appears, most likely, that this dispersal occurred over land, via the large intervening island of Mindanao. Although the exact site of collection is unknown, consideration of the collector’s itinerary suggests that it originated most likely in the vicinity of Davao City, southeastern Mindanao. Its discovery confirms that C. salitan occurs on Mindanao (or, at least, did occur on this island as recently as 133 years ago), and that locating additional populations via targeted surveys is an urgent priority for conservation of this remarkable Philippine endemic snake lineage. Keywords. Coralsnakes, distribution, Elapidae, reptiles, Southeast Asia. Introduction Conceptually, these topics invoke a number of evolutionary questions, concerning phenomena not well Compared with neighbouring Sundaland, the understood. Philippines has a relatively depauperate terrestrial elapid The first of these is the unexplained and evolutionarily snake fauna. Terrestrial Philippine snakes of the family anachronistic case of sexual size dimorphism in the Elapidae (Leviton, 1964; Leviton et al., 2014, 2018) diameter of the discharge orifice in venom-conducting include Ophiophagus (one species), Naja (three species), fangs of Naja philippinensis, which is larger in females Hemibungarus (three species), and Calliophis (four (Wüster and Thorpe, 1992). This is the only known case species). Despite the relatively low diversity of land- of this phenomenon in cobras, and such a disparity may dwelling elapid species in the archipelago, the Philippine be expected to have functional significance (Bogert, elapid snake fauna possesses interesting attributes in 1943). However, an unambiguous explanation of its terms of natural history, ecology, and biogeography. adaptive significance in Philippine cobras has yet to be provided (Wüster and Thorpe, 1992). Second, on a related point, Philippine cobras possess 1 Biodiversity Institute and Department of Ecology and highly-specialized spitter dentition (rifled, mobile fangs Evolutionary Biology, University of Kansas, Lawrence, for frontal projection of venom; Wüster and Thorpe, Kansas 66045, USA. 2 Museo Nacional de Ciencias Naturales (MNCN-CSIC), Calle 1992), and yet conflicting accounts and ambiguous José Gutiérrez Abascal 2, 28006, Madrid, Spain. reports of actual spitting behaviour in Philippine 3 Division of Biological Sciences, University of Missouri, “spitting” cobras N. philippinensis and N. samarensis Columbia, Missouri 65211, USA. (Taylor, 1922, 1975; Leviton, 1964, Leviton et al., 2014) * Corresponding author. E-mail: [email protected] suggest an unresolved disconnect between structure and © 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. function (Watt et al., 1988; Wüster and Thorpe, 1991, 1028 Rafe M. Brown et al. 1992). In addition to unclear literature accounts, we Aggregate Island Complexes, or PAICs – sensu Brown have never observed spitting behaviour in these species, and Diesmos, 2009; Brown et al., 2013a), whereas four despite numerous field encounters over the past three PAIC-endemic species of Calliophis are restricted to decades (RMB, pers. obs.). three faunal regions of the southern and western reaches Third, Philippine elapids possess unexpected and of the archipelago (Fig. 1; Leviton et al., 2018). surprising phylogenetic relationships. The close, Finally, snake biogeographers have been puzzled by sister-lineage relationship between king cobras of the the absence of kraits and large-bodied, brightly-coloured genus Ophiophagus and the Philippine endemic false coralsnakes in the archipelago. These conspicuous coralsnakes of the genus Hemibungarus was entirely faunal elements are well-documented and frequently unexpected because of their remarkably dissimilar encountered just outside of the Philippines (on the phenotypes, anatomy, ecology, microhabitats, and islands of Borneo, Sumatra, Java, Sulawesi, and the behaviour (Castoe et al., 2007; Brown et al., 2018; Thai-Malaysian Peninsula of mainland Southeast Asia; Plettenberg Laing, 2018). Uetz et al., 2020), but have been notably absent within Fourth, the biogeography of Philippine elapids is the archipelago (but see below). interesting because it involves a peculiar geographic The first four topics summarized above represent replacement pattern of coralsnakes (Leviton, 1964; reasonably well understood or, at least, partially Brown et al., 2018). In this case, the three endemic documented Philippine-specific elapid case histories that species of Hemibungarus have come to occupy three can be viewed as intriguing challenges for international separate faunal regions of the northern, northeastern, collaborations involving Philippine scientists, students, and central portions of the archipelago (the Pleistocene and researchers from nearby Southeast Asian countries Figure 1. Map of the southern Philippines with the specimen-vouchered distributions of Philippine Calliophis species indicated by various symbols as shown in the key. A yellow star marks the C. salitan type locality on northern Dinagat Island. According to the itinerary of Hipólito Fernández, collector of the first specimen ofCalliophis salitan from Mindanao Island (MNCN 22719), he accessed the island through the Port City of Davao (orange star). First Report of Calliophis salitan from Mindanao Island 1029 and beyond (Alcala, 2004a, b). However, the last point questions (Brown et al., 2018). One of these we address enumerated above has recently been addressed with the here, with the first report of a historically-significant, discovery of a strikingly distinct, spectacularly-coloured, unrecognized specimen, which the available evidence new species of long-glanded coralsnake, Calliophis suggests likely originated on the intervening island of salitan Brown et al., 2018, heretofore known from a Mindanao. The discovery of the second specimen of C. single specimen. This species, documented only from salitan, reidentified 133 years after the first record of its Dinagat Island northeast of Mindanao (Fig. 1), remains existence, and housed within the historic herpetological enigmatic because virtually nothing is known about its collections of Spain’s National Museum, the Museo biology. Calliophis salitan was discovered in 2007 and Nacional de Ciencias Naturales (MNCN), constitutes a was described on the basis of a single specimen (Brown et significant biogeographical “piece of the puzzle” in the al., 2018). Related species are referred to as “long-glanded sense that it fills a distributional gap between the central tropical coralsnakes” because they possess remarkably Philippine island of Dinagat, where C. salitan was enlarged venom glands, which extend from the head to discovered, and the Sunda Shelf landmasses of Borneo, the thoracic-cardiac cavity (Brown et al., 2018: Fig. 6). Java, Sumatra, and the Thai-Malaysian Peninsula, In the years since the initial discovery of C. salitan where its closest relatives are found (Stuebing et al., (by J.B. Fernandez in 2007) and its description (Brown 2014; Charlton, 2020). et al., 2018; Leviton et al., 2018; Weinell et al., 2019), numerous biodiversity surveys and faunal inventories Historical background and Philippine herpetological of Dinagat Island (as well as nearby Siargao Island), collections at MNCN: The “Comisión Central de southern Leyte and Samar islands, to the north, and Manila” of 1885. In his MNCN catalogue of Philippine northeastern Mindanao and Camiguin Sur islands to vertebrate specimens collected prior to 1886 and the south (Fig. 1) have been conducted by qualified assembled in 1887 as part of the Comisión’s activities, herpetologists (see Sanguila et al., 2016 for a review). Gogorza (1888) sought to summarize all that was known of However, the search for additional specimens of C. specimen-based vertebrate biodiversity of the Philippines, salitan,
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